Takhtajan Armen. Flowering Plants

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Superorder FABANAE 353

Oxyrhynchus, Phaseolus, Ramirezella, Strophostyles,

Dolichopsis, Macroptilium, Mysanthus, Oryxis; des-

modieae: Campylotropis, Kummerowia, Lespedeza,

Dendrolobium, Phyllodium, Ougeinia, Aphyllodium,

Dicerma, Ohwia, Hanslia, Arthroclianthus, Nephro-

desmus, Tadehagi, Akschindlium, Droogmansia,

Monarthrocarpus, Triﬁ dacanthus, Desmodium,

Podocarpium, Codariocalyx, Hylodesmum, Hegnera,

Pseudarthria, Pycnospora, Mecopus, Uraria,

Urariopsis, Christia, Alysicarpus, Desmodiastrum,

Melliniella, Leptodesmia, Eleiotis; psoraleeae:

Otholobium, Psoralea, Hallia, Orbexilum, Hoita,

Rupertia, Psoralidium, Pediomelum, Bituminaria,

Cullen, sesbanieae: Sesbania, Glottidium; loteae:

Hippocrepis, Scorpiurus, Securigera, Coronilla,

Podolotus, Anthyllis, Hymenocarpos, Pseudolotus,

Antopetitia, Hosackia, Ornithopus, Dorycnopsis,

Kebirita, Ottleya, Acmispon, Syrmatium, Lotus,

Dorycnium, Tetragonolobus, Tripodion, Hamma-

tolobium, Cytisopsis; robinieae: Hebsetigma,

Lennea, Gliricidia (including Yucaratonia),

Hybosema, Poitea (including Sauvallella), Sabinea,

Notodon, Corynella, Bembicidium, Olneya, Robinia,

Poissonia (including Noecracca), Coursetia, Peteria,

Genistidium, Sphinctospermum; galegeae:

Glycyrrhiza, Meristo tropis, Chesneya,

Spongiocarpella, Gueldenstaedtia, Tibetia, Erophaca,

Oxytropis, Biserrula, Astragalus (including

Neodielsia), Astracantha, Ophiocarpus, Barnebyella,

Colutea, Oreophysa, Smirnowia, Eremosparton,

Sphaerophysa, Lessertia, Sutherlandia, Swainsona,

Montigena, Clianthus, Streblorrhiza, Galega; carmi-

chaelieae: Carmichaelia, Notospar tium,

Chordospartium, Corallospartium; hedysareae:

Calophaca, Caragana, Halimodendron, Alhagi,

Eversmannia, Hedysarum, Corethrodendron, Sulla,

Taverniera, Onobrychis, Stracheya, Sartoria, Ebenus;

cicereae: Cicer; trifolieae: Parochetus, Trifolium,

Lupinaster, Amoria, Chrysaspis, Ononis, Melilotus,

Trigonella, Factorovskya, Medicago, Radiata; vicieae

(fabeae): Vicia, Lathyrus, Lens, Pisum, Vavilovia;

The most archaic genera of the family seem to be

diverse extratropical woody Caesalpinioideae –

Gleditsia, Gymnocladus, Ceratonia – Zenia, and

Cercis (Polhillet al. 1981: 2; Tucker 2002). It is there-

fore preferable to begin the system of classiﬁ cation of

Fabaceae with the subfamily Caesalpinioideae. The

most archaic tribes are Caesalpinieae, Cassieae, and

Cercideae. Partial helical order of ﬂ oral initiation is

common among Caesalpinioideae, and Gleditsia

and Ceratonia are known to have helical order through-

out organ initiation (Tucker 1991). According to

Goldblatt (1981: 455), “x = 7 would seem basic for

Caes alpinioideae, with the retention of ancestral

diploidy only in Cercis

. Polyploidy apparently became

established early in the evolution of the subfamily with

n = 14 basic in one line and n = 12 in a second more

specialized and derived line.”

Mimosoideae are closely linked to the

Caesalpinioideae. According to Elias (1981: 143), the

apparent link is between the caesalpinioid genus

Dimorphandra and the mimosoid Pentaclethra. Both

of them have similar bipinnate leaves, elongated spikes

or often paniculate inﬂ orescences, small, actinomor-

phic, bisexual ﬂ owers with imbricate sepals, ﬁ ve

fertile, alternisepalous stamens, ﬁ ve or more stamino-

dia, and pollen grains in monads. According to

Goldblatt (1981: 455), Mimosoideae are evidently of

tetraploid origin, with x = 14 and 13 frequent. In his

opinion the closest ally of Mimosoideae is probably

the Erythrophleum group of Caesalpinioideae. On

the other hand, as Polhill (1981: 191) noticed, there are

genera in Swartzieae and Sophoreae with open radial

ﬂ owers, no apparent hilar groove, and a general

similarity to some genera in the Caesalpinieae. The

less advanced position in the subfamily is occupied by

Swartzieae and Sophoreae.

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Order 89. POLYGALALES

Trees, sometimes (Xanthophyllum) up to 50 tall, or

more often shrubs, woody lianas (Balgoya up to 30 m

length), and perennial or annual herbs, very rarely

(Epirixanthes) achlorophyllous parasites; plants

unarmed, glabrous or with simple unicellular or

sometimes uniseriate hairs; bearing essential oils

or without essential oils. Xylem with tracheids (e.g.

Diclidanthera), or without tracheids. Vessels with

simple perforations; lateral pitting usually alternate.

Fibers with bordered pits. Rays heterogeneous. Axial

parenchyma mainly paratracheal, rarely

(Xanthophyllum and Moutabea) apotracheal. Sieve-

element plastids of S-type. Nodes unilacunar with one

trace. Leaves usually alternate, rarely (some Polygala)

opposite or verticillate, leathery or membranous,

sometimes reduced to scales (Salomonia and some

spp. of Polygala), simple, sometimes gland-dotted,

stipulate or estipulate, stipules intrapetiolar, free of

one another, often scaly, or spiny, or represented by

glands. Stomata anomocytic or seldom paracytic, or

anisocytic. Flowers in terminal or axillary racemes,

spikes or panicles, or rarely solitary in the axil, bracte-

ate and almost always bibracteolate, bisexual, or

(Balgoya) functionally unisexual, usually strongly

zygomorphic. Sepals ﬁ ve, imbricate free or rarely

basally connate (sometimes only lower or upper ones

connate); the two lateral (inner) sepals much larger

and petaloid. Petals free or more often adnate to sta-

mens to form a common tube, sometimes ﬁ ve, but

more often the two lateral ones much reduced or com-

pletely aborted, so that there are only three evident

petals – two upper ones and one lower median one, the

latter (carina) large, boat-shaped, and often apically

fringed. Stamens (2-) 3–10, basically ten in two cycles,

but the median member of each cycle usually sup-

pressed and the number of stamens is commonly 8, or

Superorder FABANAE 357

as a result of further reduction sometimes 7–4 or even

only 3; ﬁ laments generally more or less connate into a

cleft tube, that is more or less adnate to petals; anthers

1–2-locular, basiﬁ xed or (Xanthophyllum) more or

less dorsiﬁ xed, tetrasporangiate or disporangiate,

opening by one or two apical or subapical pores or

very short slits, or less often by long longitudinal slits

(as in Xanthophyllum), or by longitudinal valves.

Tapetum secretory. Microsporogenesis simultaneous.

Pollen grains 2–3-celled, 3-colporate, polycolporate,

often synorate. Intrastaminal nectaries present or

(Moutabeoideae) absent. Gynoecium of 2–5(-8) united

carpels; stylodia connate into a usually curved and

commonly apically 2-lobed style, one lobe stigmatic,

and the other with a tuft of hairs; ovary superior,

2-multilocular with parietal placentas and stipitate

(Xanthophyllum), sometimes pseudomonomerous (as

in Securidaca) and 1-locular, each locule with (1)2–

5(-8) or (Xanthophyllum) 4–40 ovules. Ovules

anatropous or hemitropous, epitropous, pendulous,

bitegmic, crassinucellate; micropyle zig-zag (exos-

toma often long). Female gametophyte of Polygonum-

type. Endosperm nuclear. Fruits mostly loculicidal

capsules, less often samara, nuts, drupes, or berries.

Seeds often hairy, often with a caruncle, arillate or

exarillate; testa multiplicative, exotesta subsclerotic,

endotestal cells more or less palisade or not,

U-thickened (Plisco 2000); embryo straight, sometimes

(Polygala sp.) green; cotyledons two, planoconves;

endosperm copious, oily, scanty or wanting. Contain

commonly alkaloids, ﬂ avonols (kaempferol and quer-

cetin), often saponins, commonly accumulating

aluminum, n = 5–14(-17).

Evidently related to the Fabales (see Soltis et al. 2006).

1. POLYGALACEAE

Hoffmannsegg et Link 1809 (including Diclidan-

theraceae J. Agardh 1858, Moutabeaceae Endlicher

1841, Xanthophyllaceae Gagnepain ex Reveal et

Hoogland 1990). 21/950–1000. Nearly cosmopolitan,

except for the Arctic, Polynesia, and New Zealand.

1.1 MOUTABEOIDEAE

Ovary 2–8 locular, with 1 ovule in each locule. Calyx

united with the petals into a tube. Abaxial petals not

keeled. Filaments connate into a singular tube

(Diclidanthera, Eriandra) or either in two bundles

(Moutabea) or inserted on the petals, not united into a

tube (Barnhartia). Intrastaminal nectaries absent.

Axial parenchyma apotracheal or paratracheal. – xan-

thophylleae: Xanthophyllum; mout abeae:

Balgoya, Eriandra, Barnhartia, Diclin danthera,

Moutabea.

2.2 POLYGALOIDEAE

Ovary 2–3-locular with one ovule on each axile pla-

centa. Calyx not united with the petals into a tube.

Abaxial petals keeled. Filaments united into a tube

open dorsally. Intrastaminal nectaries present. Axial

parenchyma predominantly paratracheal. – carpolo-

bieae: Atroxima, Carpolobia; poly galeae:

Bredemeyera, Acanthocladus, Badiera, Comesperma,

Epirixanthes, Muraltia (including Nylandtia),

Salomonia, Monnina, Ancy lotropis, Pseudomonnina,

Securidaca, Hualania, Pteromonnina.

Bibliography

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and D Ramos-Zamora. 1988. Morfoligia de los granos de

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Bot. Mex. 4: 21–27.

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cas acerca de 101 Polygalae americanas. Cavanillesia Altera

1: 1–456.

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Roxb. IAWA Bull. 3: 115–125.

Detienne P. 1991. Anatomic de bois de Balgoya paciﬁ ca

(Polygalaceae) de Nouvelle Caledonie. Adansonia, ser.4, 13:

17–20.

Dickison WC. 1973. Nodal and leaf anatomy of Xantophyllum

(Polygalaceae). Bot. J. Linn. Soc. 67: 103–115.

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Polygalaceae and its allied families. Agra Univ. J. Res. Sci.

11: 109–112.

Dube VP and DK Awasthi. 1985. Morphological and anatomical

studies of ﬂ oral nectaries in Polygalacaea. J. Indian Bot. Soc.

64 (2 and 3): 231–235.

Erdtman G. 1944. The systematic position of the genus

Diclidanthera Mart. Bot. Not. 1944: 80–84.

Eriksen B. 1993a. Floral anatomy and morphology in the

Polygalaceae. Plant Syst. Evol. 186: 17–32.

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nomic implications. Plant Syst. Evol. 186: 33–35.

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laceae and Valerianaceae. Thesis, Fac. Nat. Sc., Univ,.

Goteborg.

Eriksen B and C Persson. 2007. Polygalaceae. In: K Kubitzki,

ed. The families and genera of vascular plants, vol. 9,

pp. 345–363. Springer, Berlin/Heidelberg/New York.

358 Subclass VI. ROSIDAE

Leinfellner W. 1972. Zur Morphologic des Gynozeums der

Polygalaceen. Oesterr. Bot. Z. 120: 51–76.

Meijden R van der. 1982. Systematics and evolution of

Xanthophyllum (Polygalaceae). Leiden Bot. ser. 7: 1–159.

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ed. Flora Malesiana, ser. 1, 10: 455–539. Noordhoff,

Groningen.

Miller NG. 1971. The Polygalaceae in the southeastern United

States. J. Arnold Arbor. 52: 267–284.

Persson C. 2001. Phylogenetic relationships in Polygalaceae

based on plastid DNA sequences from the trn-F region.

Taxon 50: 763–779.

Plisko MA. 2000. Order Polygalales. In: A Takhtajan,

ed. Comparative seed anatomy, vol. 6, pp. 49–111. Nauka,

St. Petersburg (in Russian).

Prenner G. 2004. Floral development in Polygala myrtifolia

(Polygalaceae) and its similarities with Leguminosae. Plant

Syst. Evol. 249: 67–76.

Rao AN. 1964. An embryological study of Salomonia conton-

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Styer CH. 1937. Comparative anatomy and systematic of

Moutabeae (Polygalaceae). J. Arnold Arbor. 58: 109–145.

Verkerke W. 1984. Ovule and seed of Xanthophyllum

(Polygalaceae). Blumea 29: 409–421.

Verkerke W. 1985. Ovules and seeds of the Polygalaceae. J. Arnold

Arbor. 66: 353–394.

Verkerke W. 1991. Fruits and seeds of Balgoya paciﬁ ca

(Polygalaceae) from New Caledonia. Adansonia, ser. 4, 14:

9–12.

Westerkamp C and A Weber. 1999. Keel ﬂ owers of the

Polygalaceae and Fabaceae: a functional comparison. Bot.

J. Linn. Soc. 129: 207–221.

Superorder RUTANAE

Order 90. OXALIDALES (CONNARALES)

Small trees, shrubs, woody lianas, or, less often, herbs.

Commonly with mucilage canals and/or tanniniferous

secretory cavities. Indumentum of multicellular or

unicellular glandular hairs. Vessels with simple

perforations; lateral pitting alternate. Fibers with simple

pits, septate. Rays homogeneous or heterogeneous, all

uniseriate or occasionally biseriate for part of their

length or some of them up to ﬁ ve cells wide. Axial

parenchyma typically absent or scanty-paratracheal.

Sieve-element plastids of psc- or Pcfs-type. Nodes

trilacunar (Oxalidaceae) or pentalacunar (Connaraceae).

Leaves alternate, imparipinnate with two or more pairs

of leaﬂ ets or less often trifoliolate or unifoliolate, with

coriaceous, entire or rarely lobed, very rarely peltate

leaﬂ ets; commonly estipulate. Stomata paracytic,

anisocytic and anomo-cyclocytic. Flowers small, in

terminal or axillary panicles or racemes, bisexual or

very rarely unisexual (plants dioecious), actinomorphic

or rarely slightly zygomorphic, 5-merous or seldom

4-merous. Sepals free or rarely basally connate, imbri-

cate or valvate, usually persistent and often embracing

the base of the fruit. Petals free or rarely slightly

connate, imbricate or valvate, sometimes loriform and

circinate. Stamens ten or eight, in two cycles, the inner

sometimes staminodial; ﬁ laments slender, free, shortly

connate at the base; anthers short, dorsiﬁ xed, tetraspo-

rangiate, introrse or extrorse, opening longitudinally.

Tapetum secretory. Microsporogenesis simultaneous.

Pollen grains 2-celled, mostly 3-colporate, sometimes

3-colpate, rarely (Jollydora) 4-colpate. Nectary disc

poorly developed or wanting. Gynoecium of 5 or 1

carpels, rarely of 3, 7–8 carpels, free or only basally

connate; ovary superior; ovules anatropous, hemitrop-

ous, or more often orthotropous, bitegmic, crassinucel-

late or tenuinucellate, sometimes (Connarus, Jollydora,

and Averrhoa) with an endothelium. Female

gametophyte of Polygonum-type, or very rarely of

Allium-type. Endosperm nuclear. Fruits capsules or

indehiscent. Seeds mostly arillate, without or with

ﬁ brous exotegmen; embryo straight; endosperm

copious and oily to scanty or none; n = 5(-7)-14, 16.

Oxalidales have many similarities both with the

Rosanae (especially with Cunoniales) and with Rutanae

(especially with Sapindales).

Key to Families

1 Shrubs, woody lianas, or, less often, small trees.

Commonly with mucilage canals and/or tanninifer-

ous secretory cavities. Indumentum of multicellular

or unicellular glandular hairs. Nodes pentalacunar

with ﬁ ve traces or septalacunar with 7seven traces.

Leaves imparipinnate with two or more pairs of leaf-

lets or less often trifoliolate or unifoliolate, with

coriaceous, entire or rarely lobed, very rarely peltate

leaﬂ ets; stipules wanting. Stomata paracytic, aniso-

cytic and anomo- cyclocytic. Flowers small, in ter-

minal or axillary panicles or racemes, bisexual or

very rarely unisexual (plants dioecious), actinomor-

phic or rarely slightly zygomorphic, 5-merous or

seldom 4-merous. Sepals free or rarely basally con-

nate, imbricate or valvate, usually persistent and

often embracing the base of the fruit. Petals free or

rarely slightly connate, imbricate or valvate, sometimes

Superorder RUTANAE 359

loriform and circinate. Stamens ten or eight, in two

cycles, the inner sometimes staminodial; ﬁ laments

slender, free, shortly connate at the base; anthers

short, dorsiﬁ xed, tetrasporangiate, introrse. Pollen

grains 2-celled, mostly 3-colporate, sometimes

3-colpate, rarely (Jollydora) 4-colpate. Nectary disc

poorly developed and usually extrastaminal, or

wanting, but the receptacle sometimes nectariferous

at the center. Gynoecium of ﬁ ve or one carpels,

rarely of three, seven, or eight carpels (often some of

them abortive), free or only basally connate, often

not fully sealed, each with terminal stylodium and

capitate stigma and containing two collateral ovules

ascending from near the base, one of which may

abort; ovary superior. Ovules anatropous, hemitrop-

ous, or more often orthotropous, crassinucellate,

sometimes (Connarus, Jollydora) with an endothe-

lium. Fruiting carpels sessile or stipitate, dry,

1-seeded or sometimes (Jollydora) 2-seeded, open-

ing along the ventral suture (follicles) or seldom

along both sides (legumes), or rarely (Jollydora and

Hemandradenia) indehiscent and nutlike. Seeds

mostly arillate, exotestal with more or less devel-

oped ﬁ brous exotegmen; embryo straight, with short

hypocotyl and minute radicle; endosperm copious

and oily to scanty or none; funicle very short or

none; according to Corner (1976: 105), aril present

to some extent in all cases, primarily as an outgrowth

from the short raphe and chalaza, yellow to red,

pulpy, typically (Connarus) with a thick, often cren-

ulate, free border or short limb and a sarcotestal area

along the raphe and chalaza; with extensive free

limb (Rourea spp.); more or less wholly developed

as a sarcotesta in genera with baccate seeds. Bark,

fruits, and seeds often highly poisonous, n = 14, 16

. . . . . . . . . . . . . . . . . . . . . . . . . . . 1. connaraceae.

1 Herbs, sometimes succulent, often with ﬂ eshy rhi-

zomes or bulb-like tubers, sometimes subshrubs or

arborescent. Leaves alternate, palmately or pinnately

compound, often trifoliolate, rarely unifoliolate or

phyllodic, stipules small or absent, colleters (Oxalis)

present; leaﬂ ets usually folded back in bud and at

night. Stomata paracytic. Flowers in axillary cymes

or solitary, bisexual, usually tristylous, sometimes

cleistogamous and apetalous. Sepals ﬁ ve, basally

connate. Petals ﬁ ve, free or united at the base, con-

torted or rarely imbricate. Stamens 10 or 15, some-

times staminodal. Filaments united at the base.

Anthers extrorse. Pollen grains mostly 3-colporate,

often contain starch. Nectaries often glands opposite

the petals. Gynoecium of (3-)5 carpels, united to

form a 5-locular ovary; ovary superior, with (1)2 to

many more or less pendulous, anatropous ovules per

locule; ovules mostly tenuinucellate, rarely crassi-

nucellate and with endothelium (Averrhoa). Fruits

loculicidal, ribbed or angled capsules (Oxalis,

Biophytum) or ﬂ eshy capsules (Dapania), or

(Averrhoa, Sarcotheca) baccate. Seeds with ﬁ brous

2-layered exotegmen, endotesta walls thickened, not

palisade; testa and tegmen less differentiated when

fruit a berry; embryo large, green (Oxalis); edno-

sperm ﬂ eshy, starchy, subruminate. Present tannins,

juice acrid, with oxalated, calcium oxalatae as crys-

tals, n = (5-)7–12, mostly 7. . . . . . 2. oxalidaceae.

1. CONNARACEAE

R. Brown 1818 (including Cnestiaceae Raﬁ nesque

1830). 13/380. Pantropical, with the greatest diversiﬁ ca-

tion in Africa, Southeast Asia, and tropical America.

Cnestis, Rourea (including Bernardinia, Byrsocarpus,

Eichleria, Jaundea, Paxia, Roureopsis, Spiropetalum),

Jollydora, Connarus, Agelaea, Pseudoconnarus, Burttia,

Schellenbergia, Vismianthus, Hemandradenia, Cnestidium,

Manotes, Ellipanthus (including Pseudoellipanthus).

The family Connaraceae shares with the

Oxalidaceae. According to some botanists, including

Hallier (1912), Gundersen (1950), and Airy Shaw

(1973), Connaraceae are allied to the Oxalidaceae

(according to Hallier related to both the Leguminosae

and Oxalidaceae, according to Schellenberg (1938)

“perhaps near to Oxalidaceae,” and according to Airy

Shaw “somewhat intermediate between Leguminosae

and Averrhoaceae”). Some perhaps rather remote

afﬁ nity between Connaraceae and Oxalidaceae is sup-

ported by the same Pies-type of sieve-element plastids

(Behnke 1983).

2. OXALIDACEAE

R. Brown 1818 (including Averrhoaceae Hutchinson

1959). 5/700. Tropical and subtropical regions with a

few species widespread in temperate regions.

Oxalis, Biophytum, Averrhoa, Sarcotheca, Dapania.

360 Subclass VI. ROSIDAE

Bibliography

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Oxalidaceae: A contribution to the knowledge of P-type

plastids in dicotyledons and their signiﬁ cance. Bot. Jahrb.

Syst. 103: 1–8.

Boesewinkel FD. 1985. Development of ovule and seed coat in

Averrhoa (Oxalidaceae) with notes on some related genera.

Acta Bot. Neerl. 34: 413–424.

Boesewinkel FD and F Bouman. 2000. Oxalidaceae. In:

A Takhtajan, ed. Comparative seed anatomy, vol. 6,

pp. 21–23. Nauka, St. Petersburg (in Russian).

Breteler FJ, ed. 1989. The Connaraceae. A taxonomic study

with emphasis on Africa. Agric. Univ. Wageningen Pap.

89–6: 1–403.

Chung RCK, and AL Lim. 1998. The embryology of Averrhoa

(Oxalidaceae). Sandakania 12: 37–55.

Cocucci AA. 2004. Oxalidaceae. In: K Kubitzki, ed. The

Families and genera of vascular plants, vol. 6, pp. 285–290.

Springer, Berlin/Heidelberg/New York.

Devi DR and LL Narayana 1990. Systematic position of

Averrhoa (Oxalidaceae). Feddes Repert. 101: 165–170.

Dickison WC. 1971–1974. Anatomical studies in the

Connaraceae: I. Carpels. II. Wood anatomy. III. Leaf

anatomy. IV. The bark and young stem. J. Elisha Mitchell

Sci. Soc. 87: 77–86, 1971; 88: 120–136; 89: 121–138, 1973;

166–171, 1973 (1974).

Dickison WC. 1979. A survey of pollen morphology of the

Connaraceae. Pollen Spores 21: 31–79.

Forero E. 1983. Connaraceae. Flora Neotrop. Monograph 36.

New York.

Heimsch C. 1942. Comparative anatomy of the secondary xylem

in the “Gruinales” and “Terebinthales” of Wettstein with ref-

erence to taxonomic grouping. Lilloa 8: 83–198.

Huynh K-L. 1969. Étude du pollen des Oxalidaceae Morphologic

générale — Palynotaxonomie des Oxalis americains. Bot.

Jahrb. Syst. 89: 272–303.

Jongkind CCH, and RHJ Lemmens. 1989. The Connaraceae:

A taxonomic study with special emphasis on Africa.

Landbouwuniversiteit te Wageningen.

Knuth R. 1930. Oxalidaceae. In: A Engler, ed. Pﬂ anzenreich, 95

(IV. 130): 1–481. Berlin.

Leenhouts PW. 1958. Connaraceae. In: CGGJ van Steenis ed.

Flora Malesiana ser. 1, 1 (5): 495–541. Noordhoff, Groningen.

Leinfellner W. 1970. Über die Karpelle der Connaraceen.

Oesterr. Bot. Z. 118: 542–559.

Lemmens RHMJ, EJ Breteler, and CCH Jongkind. 2004.

Connaraceae. In: K Kubitzki, ed. The families and genera of

vascular plants, vol. 6, pp. 74–81. Springer, Berlin/

Heidelberg/New York.

Matthews ML and PK Endress. 2002. Comparative ﬂ oral mor-

phology and systematics in Oxalidales (Oxalidaceae,

Connaraceae, Brunelliaceae, Cephalotaceae, Cunoniaceae,

Elaeocarpaceae, Tremandraceae). Bot. J. Linn. Soc. 140:

321–381.

Mosebach G. 1934. Die Fruchtstielschwellung der Oxalidaceen

und Geraniaceen. Jahrb. Wiss. Bot. 79: 353–384.

Mondal MS. 1990. Pollen morphology and systematic relation-

ships of families Sabiaceae (s.l.) and Connaraceae. Botanical

Survey of India, New Delhi.

Narayana LL. 1966. A contribution to the ﬂ oral anatomy of

Oxalidaceae. J. Jpn. Bot. 41: 321–328.

Narayana LL. 1970. Oxalidaceae, Geraniaceae. In Symposium

on comparative embryology of angiosperms. Bull. Indian

Natl. Sci. Acad. 41: 114–120.

Oltmann O. 1971. Pollenmorphologiscyh-systematische

Untersuchungen innerhalb der Geraniales. Diss. bot. 11, 163

+ XI pp. Cramer, Lehre.

Radlkofer L. 1886. Über die dursichtigen Punkte und andere

anatomische Charakter der Connaraceae. Sitzungsber. Bayer.

Akad. Wiss. (M.-Ph. Kl.) 16: 345–378.

Rama Devi D. 1991. Floral anatomy of Hypseocharis

(Oxalidaceae) with a discussion on its systematic position.

Plant Syst. Evol. 177: 161–164.

Reddy B Bal and LL Narayana. 1982. Systematic position of

Averrhoaceae. J. Econ. Taxon. Bot. 3: 343–348.

Robertson KR. 1975. The Oxalidaceae in the southeastern

United States. J. Arnold Arb. 56: 223–239.

Schellenberg G. 1925. Die phylogenetische Entwiclung und die

Wanderungen der Connaraceen. Bot. Jahrb. Syst. 60:

207–251.

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Comparative seed anatomy, vol. 5, pp. 299–311. Nauka,

St. Petersburg (in Russian).

Order 91. SAPINDALES

Trees, shrubs, woody lianas, or sometimes subshrubs or

herbs. Vessels with simple perforations or less often

perforations scalariform; lateral pitting alternate or

sometimes opposite. Fibers with bordered or simple pits,

often septate. Rays heterogeneous or homogeneous.

Axial parenchyma mostly paratracheal, sometimes

scanty or even wanting. Sieve-element plastids of

Ss-type. Nodes usually trilacunar. Leaves opposite or

less often alternate, simple or more often compound,

estipulate or sometimes with small, caducous stipules.

Stomata of diverse types. Flowers in various types of

inﬂ orescences, rarely solitary, bisexual or more often

unisexual, actinomorphic or less often more or less zygo-

morphic (strongly zygomorphic in Hippocastanaceae),

usually 5–4-merous. Sepals free or more or less connate,

imbricate or rarely (some Sapindaceae) valvate. Petals

free, equal or unequal, imbricate or rarely valvate (some

Sapindaceae) or seldom wanting. Nectary disc of recep-

tacular origin commonly present, extrastaminal or less

often (Aceraceae) intrastaminal. Stamens (4)5-10, rarely

more; ﬁ laments free; anthers tetrasporangiate, introrse,

basiﬁ xed or more often dorsiﬁ xed, opening longitudi-

nally. Tapetum secretory. Microsporogenesis simultane-

ous. Pollen grains 2-celled or 3-celled mostly 3-colporate.

Gynoecium of 3-2(4-5) united carpels; stylodia free or

Superorder RUTANAE 361

more or less connate; ovary superior, with (1)2(-4-10)

ovules per locule. Ovules epitropous or apotropous,

anatropous or less often hemitropous, campylotropous

or orthotropous, bitegmic, crassinucellate. Female

gametophyte of Polygonum-type. Endosperm nuclear.

Fruits of various types. Seeds exotestal, mesotestal, or

rarely (Alectryon in Sapindaceae) with ﬁ brous

exotegmen; embryo mostly curved; endosperm copious,

or very scanty or lacking.

The order Sapindales most probably originated from

some cunonialean ancestor. The Staphyleaceae, which

are relatively the least advanced member of the order

(Takhtajan 1959, 1966, 1987; Cronquist 1981, 1988),

have much in common with the Cunoniaceae (and

related families) that Hallier (1908, 1912) placed these

two families side by side in his order Rosales sensu

lato, in which he was later followed by van Steenis

(1959). The Staphyleaceae are undoubtedly so closely

related to Cunoniaceae and related families (especially

to the Brunelliaceae) that there is not much objection to

their inclusion in the order Cunoniales. However, it

seems to me that the similarities between Staphyleaceae

and the sapindalean families are greater than those

between Staphyleaceae and Cunoniales. “It is interest-

ing and perhaps signiﬁ cant that the Staphyleaceae have

the same basic chromosome number (13) as the

Aceraceae,” says Cronquist (1981:791). “The charac-

teristic inﬂ ated capsules of Staphylea are much like

those of Koelreuteria in the Sapindaceae.” In many

respects, Staphyleaceae represent a link between the

Sapindales and Cunoniales. However, as Doweld (1996)

states, exotegmic seeds of Cunoniaceae are more spe-

cialized, than exo-mesotestal seeds of Staphyleaceae. It

is therefore more probable, that the Staphyleaceae

derived not directly from the Cunoniaceae, but rather

from their less specialized ancestor.

Key to Families

1 Vessels with scalariform perforation.

2 Intrastaminal nectary disc well developed. Leaves

stipulate. Seeds with copious endosperm. Trees

and shrubs, evergreen or deciduous, non-laticif-

erous and without coloured juice; wood often

ﬂ uorescing; ﬁ bers with bordered pits; vessels

with scalariform perforation. Druses and frothy

mucilage cells widespread in Dalrympelea. Axial

parenchyma scanty paratracheal. Nodes pentala-

cunar with ﬁ ve traces. Leaves opposite, with

well-developed caducous stipules, imparipinnate

or often trifoliolate, occasionally (some species

of Staphylea) unifoliolate, serrate or dentate;

stipules usually present, but sometimes reduced

to glands or absent. Stomata anisocytic. Flowers

in racemes or in panicles, bisexual or unisexual

(plant monoecious, dioecious, or polygamo-

monoecious), small, actinomorphic, 5- merous.

Sepals more or less free, imbricate; petals imbri-

cate. Stamens ﬁ ve, isomerous with the perianth;

ﬁ laments glabrous to pubescent; anthers dorsi-

ﬁ xed, introrse, tetrasporangiate. Pollen grains

3-colporate or rugate. Carpels 2-3(-4), almost

free or partly connate. Ovary superior to semi-

inferior, laterally and sometimes also apically

lobed, with (1-)6-12 apotropous ovules in two

rows in each locule. Fruits multifollicles, inﬂ ated

capsules opening at the tip, or berries

(Dalrympelea). Seeds arillate or exarillate, exo-

mesotestal; chalaza nonbullate; radicle short;

embryo straight, green; endosperm scanty

(Staphylea japonica) or ﬂ eshy; cotyledons large.

Present cyanidin, kaempferol and quercetin, n =

11, 13, 14. . . . . . . . . . . . . . . . 1. staphyleaceae.

2 Intrastaminal nectary disc poorly developed or

wanting. Evergreen trees with reddish wood

(Huertea) or (Tapiscia) deciduous trees; the wood

without growth rings. The imperforathe tracheary

elements septate ﬁ bre-tracheids with reduced bor-

ders on the pits, or septate libriform ﬁ bers with

simple pits. Nodes trilacunar. Leaves 2–10-jugate;

leaﬂ ets sometimes with pocket-shaped domatia or

with hairy domation in axils of and along lateral

nerves; bases of petioles, petioluli and blades of

the leaﬂ ets bearing several small glands. Stipules

lacking (Huertea) or reduced to glands (Tapiscia).

Stomata anisocytic. Flowers 5(6)-merous; sepals

fused only in low third, copular. Petals covered

with multicellular trichomes. Anthers dorsiﬁ xed,

with (Tapiscia) or without a ﬁ brous endothecium.

Pollen grains rugate or polyrugate. Nectary disc

poorly developed, 5-lobed, or lacking (Tapiscia).

Ovary 1-locular (Tapiscia), style elongate,

unbranched, stigma bilobed, or (Huertea) basally

2-locular, with two spreading stylar branches;

ovules 1–2 in each locule, sessile or attached to

an axile placenta. Fruit 1-seeded, without sclero-

tis tissue, apparently indehiscent (Tapiscia) or

baccate (Huertea), with thin, ﬂ eshy or crusta-

ceous pericarp and without sclerotic tissue. Seeds

362 Subclass VI. ROSIDAE

of Huertea subglobose, brownish, smooth, with

sclerotic cells in the mesotesta; seed-coat rather

thin; vascular bundle spreading into ﬁ ne rows of

tracheids in the bullate chalaza, separated from

the endosperm; raphe with hard sclerotic tissue

of ligniﬁ ed pitted cells; endosperm thick-walled,

oily; embryo with rather long radicle; cotyle-

dons ﬂ at; seeds of Tapiscia plump, micropype

acute at the base of the loculus, the adjacent

hilum small, chalazal area slightly depressed;

endosperm thin-walled, horny, oily; embryo

small with relatively long radicle and small thin

cotyledons. . . . . . . . . . . . . . . .2. tapisciaceae.

1 Vessels with simple perforation.

3 Leaves alternate or very rarely opposite, often

gland-dotted, often pulvinate; usually ternate,

or pinnate, or bipinnate, or multiply compound

( sometimes biternate), seldom simple; stipules

wanting in trees, frequently present in lianas,

sometimes conspicuously large. Trees and

shrubs, sometimes woody or rarely herba-

ceous (Cardiospermum) lianas, these often

with tendrils, which are transformed inﬂ ores-

cence axes, and often their stems with unusual,

secondary anomalous vascular structure.

Vegetative organs usually contain resinous or

latex-like secretions in special cells. Fibers

with simple pits, commonly septate. Rays het-

erogeneous. Axial parenchyma paratracheal,

or apotracheal and paratracheal. Nodes

trilacunar. Stomata anomocytic or sometimes

paracytic. Flowers small, in terminal or axil-

lary cymose inﬂ orescences, rarely solitary and

axillary, sometimes cauliﬂ orous in lianas, usu-

ally unisexual (plants monoecious or polyga-

momonoecious), actinomorphic or more often

obliquely zygomorphic. Sepals 5(4), free or

sometimes basally connate, usually imbricate,

sometimes valvate. Petals mostly 5(4), imbri-

cate, sometimes more than 5 or only 3 or want-

ing, often with basal scalelike appendage

concealing a nectary. Nectary disc usually

well marked, annular or often unilateral,

extrastaminal or rarely (Dodonaea) intrastami-

nal and minute. Stamens 4-5(-6), or 8, or 10,

or 11–100; ﬁ laments free, often hairy. Anthers

dorsiﬁ xed, or basiﬁ xed, to slightly ventriﬁ xed,

more or less versatile, usually apically append-

aged, opening longitudinally. Pollen grains

2-celled, usually 3-colporate, sometimes

2-colporate or syncolporate, or rarely

3–4-porate. Gynoecium of (2-)3(-8) united

carpels; stylodia more or less free or united

into a terminal, often lobed or cleft style;

stigma simple or lobed; placentation usually

axile, sometimes parietal; ovary usually 3-loc-

ular, superior, locules without ‘false septa’,

usually with one ovule, less often

(Dodonaeoideae) with two or several ovules

per locule. Ovules pendulous or horizontal, or

ascending, anatropous to hemitropous or

campylotropous, often without a denned funi-

cle, but broadly attached to placental protuber-

ance. Fruits capsules, drupes, schizocarps, or

berries, often red, sometimes winged (Serjania,

Thouinia, Paullinia, and Diatenopteryx); cap-

sules septifragal or loculicidal (Cupania).

Seeds often arillate or with sarcotesta (in

dehiscent fruits), usually with curved oily and

starchy, often green embryo, often with plicate

or twisted cotyledons; endosperm lacking.

Present proanthocyanidins, ﬂ avonols, toxic

saponins, cyclopropane amino acids, some-

times alkaloids, n = 10–16.. .3. sapindaceae.

3 Leaves opposite.

4 Flowers actinomorphic, small, some or all

functionally or truly unisexual (andromonoe-

cious, androdioecious, or polygamodioecious),

in terminal or sometimes axillary racemes

sometimes contracted to corymbs or umbels,

rarely in large panicles. Sepals ﬁ ve or rarely

four, free or rarely basally connate, imbricate.

Petals ﬁ ve, rarely four or wanting, free, often

sepal-like, shortly clawed. Nectary disc usually

present, extrastaminal, usually ﬂ at but some-

times lobed or divided or reduced to teeth,

rarely absent. Stamens 4-10(12), mostly 8;

anthers slightly dorsiﬁ xed or basiﬁ xed, versa-

tile. Pollen grains mostly 3-colpate or 3-colpo-

rate. Vestigial gynoecium often present in male

ﬂ owers. Gynoecium of two carpels, stylodia

divergent and free or basally connate, each with

a terminal stigma; ovary 2-locular, usually

compressed at right angels to the septum, with

two axile ovules in each locule. Ovules pendu-

lous, anatropous to almost orthotropous. Fruits

compressed, splitting when ripe into two usu-

ally 1-seeded, nutlike, winged mericarps, sep-